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Systematic Parasitology 54: 81–88, 2003.

© 2003 Kluwer Academic Publishers. Printed in the Netherlands.


81

Viscachataenia n. g., with the redescription of V. quadrata (von Linstow,


1904) n. comb. (Cestoda: Anoplocephalidae) in Lagidium viscacia
(Rodentia: Chinchillidae) from Argentina

G. Denegri1,2 , M. C. Dopchiz2 , M. C. Elissondo1,2 & I. Beveridge3


1 Departamento de Biologia, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata,
Funes 3250, Mar del Plata, Buenos Aires, Argentina
2 Consejo Nacional de Investigationes Cientı́ficas y Técnicas, Ciudad de Buenos Aires Argentina
3 Department of Veterinary Science, University of Melbourne, Parkville, Victoria, 3052, Australia

Accepted for publication 19th July, 2002

Abstract
A new genus, Viscachataenia, is erected to accommodate the anoplocephalid cestode Cittotaenia quadrata von
Linstow, 1904, based on a redescription of the species from material collected from the chinchillid rodent Lagidium
viscacia in Argentina. The new genus is characterised by paired genitalia, a reticulate uterus and the vagina entering
the genital atrium anterior to the cirrus-sac. Viscachataenia therefore has similarities with Monoecocestus Beddard,
1914, a genus which is common in South American rodents but which has a single set of genitalia in each segment.
Cittotaenia viscaciae (Spasskii, 1951) and Bertiella findlayi Mazza, Parodi & Fiora, 1932, also from viscachas, are
considered synonyms of V. quadrata.

Introduction M. pectinata as its type-species, and also placed in it


the specimen described by Joyeux & Dollfus (1931) as
Two species of anoplocephalid cestodes with paired a new species, M. viscaciae.
genitalia have been described from viscachas belong- Beveridge (1978) redescribed M. viscaciae from
ing to Lagidium Meyen (Rodentia: Chinchillidae) the single incomplete specimen, showing that it did not
from South America. von Linstow (1904) provided belong within Mosgovoyia, as it had a reticulate uterus
a brief description of Cittotaenia quadrata, collected and was therefore more similar to Cittotaenia Riehm,
from Lagidium peruanum Meyen in Peru, based on a 1881 to which it was transferred. Beveridge (1978)
specimen sent to him by Sir Arthur Shipley in Cam- also examined the extant type-material of C. quadrata
bridge. No additional details associated with the col- and noted a number of similarities with C. viscaciae.
lection were provided. The species name was applied He concluded however that, although C. quadrata and
because of the rather characteristic egg, which was C. viscaciae might be the same species, the existing
four-lobed (von Linstow, 1904). Subsequently, Joyeux specimens did not allow any firm conclusions to be
& Dollfus (1931) reported a specimen of Cittotaenia drawn.
pectinata (Goeze, 1782) probably from L. viscacia The other species of anoplocephalid cestodes
from Valdivia, Chile, held in the collections of the known from viscachas have a single set of genitalia in
Munich Museum. They noted that the species dif- each segment. Mazza et al. (1932) described Bertiella
fered somewhat from C. pectinata, a species found finleyi Mazza, Parodi & Fiora, 1932 from C. viscaciae
normally in leporids (Beveridge, 1978), but consid- in Argentina, a species which has not been reported
ered that C. pectinata was a morphologically variable since its original description, while Parra (1953) re-
species based on Baer’s review (1927) of the fam- ported Monoecocestus thelkeldi (Parra, 1952) from
ily Anoplocephalidae and allocated their specimen to Lagidium peruanum in Peru.
this species. Spasskii (1951) erected Mosgovoyia, with
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The collection of new material from viscachas large. Segments craspedote, wider than long. Geni-
from Argentina has allowed a re-examination of this talia paired. Genital ducts cross osmoregulatory canals
problem. In the present paper, C. quadrata is re- dorsally. Cirrus-sac powerfully developed; internal
described from new material and C. viscaciae is placed seminal vesicle present; external seminal vesicle ab-
tentatively as a synonym of C. quadrata. Bertiella sent. Testes arranged in two laterally-extended groups
finleyi is in part a synonym of C. quadrata. A new on posterior border of segment. Ovaries poral. Vagina
genus is erected to accommodate C. quadrata, be- enters genital atrium anterior to cirrus-sac. Seminal
ing allied with Monoecocestus Beddard, 1914, which receptacle absent. Uterus initially transverse, tubular,
occurs commonly in South American rodents (Rego, becoming reticulate, then saccate. Pyriform apparatus
1961). present. Parasitic in chinchillid rodents; South Amer-
ica. Type-species: V. quadrata (von Linstow, 1904) n.
comb.
Materials and methods
Viscachataenia quadrata (von Linstow, 1904) n.
Fourteen specimens of Lagidium viscacia boxi comb.
Thomas were collected from Las Bayas, Ñorquincó Syns Cittotaenia quadrata von Linstow, 1904;
Department, Río Negro Province, Argentina. Details C. pectinata (Goeze, 1782) sensu Joyeux & Dollfus
of the collection methods and locality are provided by (1931); Mosgovoyia viscaciae Spasskii, 1951; C. vis-
Led et al. (1979). Cestodes were washed in running caciae (Spasskii, 1951) Beveridge, 1978; Bertiella
water, fixed in 5% formaldehyde and subsequently finlayi Mazza, Parodi & Fiora, 1932 (in part)
stored in 70% ethanol. Five cestodes were stained
with Celestine blue, dehydrated in ethanol and cleared Specimens examined
in methyl salicylate. The outer muscle layers were Type-specimen of Cittotaenia quadrata von Linstow,
removed using fine forceps and cestodes were subse- 1904, 8 slides of serial sections, 5 frontal, 3 transverse,
quently mounted in Canada balsam. Mature regions of MNHG 126/64-71.
cestodes were embedded in paraffin, and serial trans- Type-specimen of Cittotaenia viscaciae (Spasskii,
verse sections cut at a thickness of 5 µm were stained 1951), 12 slides of serial sections, MNHG 126/16-27.
with haematoxylin and eosin. Gravid segments were Five specimens from Lagidium viscacia boxi, Las
teased apart in water to release eggs and, following Bayas, Argentina, MLPA 5066, MHNG INVE 33493-
examination in water, eggs were cleared in lactophenol 4.
and subsequently mounted permanently in polyvinyl
lactophenol. Coverslip pressure applied to some eggs Description (Figures 1-12)
resulted in expulsion of the pyriform apparatus, allow-
ing examination of its morphology and surrounding Large cestodes, 185-320 (250) long, maximum width
membranes. 10. Scolex 0.40-0.53 (0.39, n=5) in diameter, lacking
Drawings were made with the aid of a drawing rostellum (Figure 1); 4 sub-circular suckers, 0.13-0.17
tube attached to an Olympus BH microscope. Mea- (0.16) × 0.12-0.17 (0.16). Neck absent. Segments
surements were made using an ocular micrometer and transversely elongate, prominently craspedote. Im-
are presented in millimetres, unless otherwise speci- mature segments much wider than long, 0.32-1.00
fied, as the range of 10 measurements followed by the (0.56, n=5) × 3.6-6.5 (5.2, n=5). Mature segments
mean in parentheses. 0.35-1.10 (0.75) × 4.4-7.1 (5.7).
Specimens have been deposited in the Museo Genitalia paired. Genital atrium in middle of lat-
de la Plata (MLPA), Argentina and in the Muséum eral margin of segment, deep, with thick corrugated
d’Histoire Naturelle, Geneva (MHNG). walls. Cirrus-sac robust, with thick muscular walls,
broadest near distal extremity (Figure 4), 0.67-0.90
(0.77) × 0.22-0.39 (0.31) in mature segments, 0.70-
Viscachataenia n. g. 1.10 (0.91) × 0.34-0.39 (0.37) in gravid segments;
distal cirrus straight, thick walled, armed with spines
Diagnosis 20 µm long (Figures 2, 4); proximal cirrus slightly
Anoplocephalidae Cholodkovsky, 1902. Strobila sinuous, with few spines; internal seminal vesicle
subcircular, 0.16-0.25 (0.20) × 0.10-0.19 (0.15) in
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mature segments, 0.41-0.52 (0.46) × 0.21-0.32 (0.26) nal subdivisions, filling medulla and extending lateral
in gravid segments. When everted, genital atrium and dorsal to osmoregulatory canals (Figure 12). Egg
forms conical projection or genital papilla at base of thick-shelled, generally 4-lobed in shape, with size of
cirrus; internal seminal vesicle not evident in everted individial lobes varying considerably (Figures 6, 7);
cirri (Figure 3). Strong retractor muscle leads medially some eggs 3-lobed (Figure 8). External surface of eggs
from proximal pole of cirrus-sac (Figure 4). Exter- with filamentous projections arranged in 4, or more
nal seminal vesicle absent; vas deferens coiled, passes rarely 3 groups, depending upon number of lobes to
posteriorly and medially towards testes; in vicinity of egg (Figures 6-8). Oncosphere 15-19 (17)µm in di-
testes, vas deferens branches, giving off fine vasa ef- ameter, surrounded by elongate pyriform apparatus,
ferentia to individual testes. Testes small, spherical, 33-38 (35) × 18-23 (20)µm; distal extremity of pyri-
0.03-0.05 (0.04) in diameter, distributed in 3-4 dorso- form apparatus attached to inner envelope by strands
ventral layers on dorsal aspect of medulla (Figure 5); of tissue, sometimes suggesting presence of elongate
testes arranged in 2 groups per segment, with 80-107 horns extending from pyriform apparatus, which in
(90) testes per group. Testes extend from vitellarium fact represent foldings of inner envelope.
medially towards mid-line of segment; medial termi- Osmoregulatory system consists of simple ring an-
nation of testis groups variable, with some individual terior to suckers (Figure 1) into which paired dorsal
testes or small groups of testes scattered between and ventral canals empty; dorsal canals narrower, ex-
major fields; considerable variation occurs between ternal to ventral canals, diameter 0.03-0.07 (0.06);
adjacent segments (Figure 9). ventral canals medial to dorsal canals, wider, 0.06-
Vagina simple, tubular, entering genital atrium an- 0.16 (0.13) in diameter; transverse canal 0.07 in di-
terior to cirrus-sac (Figure 4). Vagina runs initially ameter, connects ventral canals at posterior margin of
along anterior margin of cirrus-sac, then runs poste- each segment. Musculature strongly developed. Lon-
riorly and medially, terminating between ovary and gitudinal musculature consists of 2 layers of fibre
vitellarium; proximal vagina variously distended by bundles, each with up to 10 fibres per bundle; trans-
sperm (Figures 4, 5), not forming distinctive semi- verse muscles present in band internal to longitudinal
nal receptacle. Ovary flabelliform, on ventral aspect musculature; dorso-ventral muscles well developed,
of medulla, 0.23-0.45 (0.36) × 0.35-0.45 (0.41), with individual fibres cross medulla at irregular intervals.
numerous clavate lobules radiating anteriorly and pos-
teriorly from central oöcapt. Vitellarium reniform,
slightly lobulate, dorsal and slightly poral to ovary, at Discussion
level of oöcapt, 0.14-0.19 (0.16) × 0.17-0.22 (0.20).
Short fertilisation canal joins vitelline duct immedi- Although von Linstow’s description of C. quadrata is
ately before entry to Mehlis’ gland. Mehlis’ gland brief, the cestodes described above conform with his
anterior to vitellariun, not visible in most whole- description in most respects. In particular, the eggs are
mounts, subcircular, 90 µm in diameter. Uterine duct four-lobed, apparently a characteristic of the species
leads dorsally from Mehlis’ gland (Figure 5), then (von Linstow, 1904). In addition, the specimens agree
turns ventrally and runs anteriorly to uterus. Uterus with all significant features visible in the type-slides of
initially a single transverse cord extending from os- C. quadrata in the MNHG. A minor difference which
moregulatory canals, between ovary and vitellarium, exists is the description and illustration of a seminal
then medially, ventral to testes (Figure 5). Uterus sub- receptacle by von Linstow (1904). In the new material,
sequently develops dorsal and ventral branches from the proximal end of the vagina is variously dilated, but
median tube, eventually forming reticulated network as the shape of the dilatation is not consistent, it has
extending across segment, except at distal extremi- not been identified as a seminal receptacle. von Lin-
ties where broad tubular extensions develop, extending stow (1904) illustrated (Figure 3) the vagina emptying
to but not beyond osmoregulatory canals (Figure 10). into the genital atrium dorsal to the cirrus-sac. This
Subsequent uterine development involves expansion situation could not be confirmed in the type-slides and
of tubular network, with most posterior part of uterus is possibly an error. The position of the ovary was
consisting of single transverse tube from which other described by von Linstow (1904) and illustrated (Fig-
reticulations extend; uterus confined within osmoregu- ure 3) as dorsal to the vitellarium and Mehlis’ gland.
latory canals. Gravid segments 0.70-1.10 (0.91) × 5.6- Spasskii (1951) commented that such an arrangement
10.3 (8.5). Gravid uterus saccate, with numerous inter- was most unlikely and is almost certainly an error.
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Figures 1-8. Viscachataenia quadrata (von Linstow, 1904) n. g., n. comb. from Lagidium viscacia. 1. Scolex. 2. Histological section through
distal part of cirrus-sac showing vagina anterior to cirrus-sac. 3. Everted cirrus showing peduncle formed by everted genital atrium and lack
of obvious internal seminal vesicle. 4. Genitalia, dorso-ventral view. 5. Transverse histological section through genitalia in mature segment,
dorsal aspect towards top of page. 6-8. Eggs, showing variation in shape and number of lobes of egg shell. Scale-bars: 1-5, 0.1 mm; 6-8,
0.01 mm. Abbreviations: c. cirrus; do, dorsal osmoregulatory canal; isv, internal seminal vesicle; m, Mehlis’ gland; o, ovary; t, testis; u, uterus;
v, vitellarium; va, vagina; vd, vas deferens; vo, ventral osmoregularory canal.
Figures 9-12. Viscachataenia quadrata (von Linstow, 1904) n. g., n. comb. from Lagidium viscacia. 9. Mature segments showing variation in the distribution of testes. 10. Post-mature segment
showing reticulate uterus which forms following the dissolution of the female genitalia; the testes persist. 11. Pre-gravid segment in which the uterus is filling with eggs, losing its reticulate
appearance, but is confined within the osmoregulatory canals. 12. Gravid segment with fully developed uterus obscuring the osmoregulatory canals. Scale-bars: 0.1mm.
85
86

In the new material, these structures were found in closely related species of cestodes in Lagidium, but
their ususal dorso-ventral relationships. Despite these the evidence at present is not substantive.
differences, which are probably simple errors in von Mazza et al. (1932) described a new species of
Linstow’s (1904) account, the cestodes described here anoplocephalid cestode, Bertiella finlayi, collected
are considered to be C. quadrata. from L. viscacia ( as L. tucumanus Thomas) in Jujuy
C. viscaciae is treated tentatively as a synonym of Province, Argentina. They described collecting both
C. quadrata following comparison of the new speci- adult and juvenile cestodes. Attempts to locate type-
mens with the type-slides of C. viscaciae. Beveridge specimens were unsuccessful. However, examination
(1978) noted that the structure of the cirrus-sac was of their published figures and photographs suggests
similar in the two species, and suggested that they that while the adult specimens they reported were
might be synonyms. However, because of the poor probably C. quadrata, the ‘juvenile’ specimens were
quality of the type-material of C. quadrata and the a related cestode, Monoecocestus threlkeldi (Parra,
incomplete nature of the type-specimen of C. visca- 1953). Figures 2-4 of Mazza et al. (1932) are of ma-
ciae, Beveridge (1978) concluded that new material ture and gravid specimens of a small cestode which
was required before any judgement could be made on is indistinguishable from the original description of
the matter. The current description agrees with that of M. thelkeldi given by Parra (1953). The large cestodes
Beveridge (1978) in most respects. Beveridge (1978) they reported are of a similar size to C. quadrata, and
stated that an internal seminal vesicle was absent, but although Mazza et al. (1932) gave a description of
in the new material it is evident that the vesicle may the internal morphology of the segments which cor-
be slender or inapparent in some segments, partic- responds closely with M. threlkeldi, they did include
ularly when the cirrus is everted. Beveridge (1978) drawings and photographs of the eggs, most of which
also described a seminal receptacle but, as explained are four-lobed and have groups of filamentous pro-
above, in the new material, the structure is quite vari- jections on the shell. Their figure 9, of a five-lobed
able in shape. The egg of C. viscaciae was described egg, represents a feature not seen in the material de-
by Beveridge (1978) as spherical rather than four- scribed above. There seems little doubt therefore that
lobed as is the case in C. quadrata, a difference which the large cestodes described by Mazza et al. (1932),
is not readily explained, but could be related to the despite of the absence of types, are C. quadrata. For
degree of maturity of the eggs. The eggs of C. visca- this reason, B. finlayi has been included as a synonym
ciae are observable only in utero, and in comparable of C. quadrata.
eggs of C. quadrata, while the lobed nature of the The development of the uterus of C. quadrata has
egg is evident, the fine, hair-like projections on the not been described previously. von Linstow (1904) de-
surface of the egg are not visible. Beveridge (1978) scribed it simply as filling the segment, with dorsal and
also described the vagina entering the genital atrium ventral diverticula. Beveridge (1978), when describ-
posterior to the cirrus-sac. However, this is shown in ing C. viscaciae, noted that the uterus was reticulate
only one figure (Beveridge, 1978, figure 71) in which and, on these grounds, removed the species from Mos-
the posterior margin of the genital papilla, formed by govoyia which has a simple tubular uterus (Spasskii,
evagination of the genital atrium, may have been mis- 1951; Beveridge, 1978), placing it tentatively in Cit-
taken for the termination of the vagina. In the new totaenia, the type-species of which, C. denticulata
material, the position of the distal vagina is extremely (Rudolphi, 1904), has a slightly reticulate uterus.
difficult to see in most segments and is only clearly The uterus of C. quadrata forms initially as a sin-
visible in a few segments in which the cirrus-sac has gle transverse cord and subsequently develops dorsal
been displaced while removing the outer muscles with and ventral branches, producing a complex reticulate
forceps. In sections, the vagina is readily detectable as uterus stretching across the segment, with a series of
a very slender tube anterior to the cirrus-sac (Figure 2). parallel branches extending laterally at each extremity.
The difficulty of tracing the distal vagina probably ac- The fully-developed uterus is saccate. This form of
counts for the conflicting descriptions of von Linstow development differs from those described by Spasskii
(1904) and Beveridge (1978). C. viscaciae is therefore (1951) and Rausch (1976), but resembles the form of
treated provisionally as a synonym of C. quadrata. uterine development found in Paranoplocephala om-
It is possible that the differences in the shape of the phalodes (Hermann, 1783), as described by Rausch
egg indicate the existence of two distinct but very (1976), in which the uterus forms as a transverse cord-
like aggregation of cells, developing into a transverse
87

band which becomes fenestrated and eventually fills The original host of V. quadrata was identified by
the segment as a sac with diverticula. It is likely von Linstow (1904 ) as Lagidium peruanum, a species
that, as uterine development is described in detail in occurring mainly in Peru (Nowak, 1999). The host
a greater number of species, more variations on the of C. viscaciae is uncertain (Beveridge, 1978), but is
currently accepted types of uterine development will probably L. viscacia, which occurs in Chile (including
be recorded. the area around Valdivia), Peru, Bolivia and Argentina
The presence of a reticulate uterus and paired geni- (Nowak, 1999). The current report confirms the occur-
talia indicates similarities with Cittotaenia, Moniezia, rence of the species in L. viscacia, although L.v. boxi
Blanchard, 1891 and Diandrya Darrah, 1930. How- is considered by some authors (Rowlands, 1974) to
ever, C. quadrata differs from all of these genera in represent a distinct species. Nowak (1999) followed
having the vagina enter the genital atrium anterior to Cabrera (1961) in recognising only three species of
the cirrus-sac, a characteristic observed only in Mo- Lagidium, L. peruanum, L. viscacia and L. wolffsohni
noecocestus, a genus which has a single set of genitalia Thomas, the latter species occurring in the extreme
per segment. The unusual egg of C. quadrata, with south of Chile and Argentina. The opinions of Cabrera
groups of fine hair like projections, has parallels in (1961) and Nowaks (1999) are followed here, suggest-
species of Monoecocestus with the eggs of M. amer- ing that V. quadrata occurs in both L. peruanum and
icanus (Stiles, 1895), M. thomasi Rausch & Maser, L. viscaciae.
1977 and M. variabilis (Douthitt, 1915) which also
have their surfaces uniformly covered with fine projec-
tions (Freeman, 1949; Rausch & Maser, 1977). Other Acknowledgement
species in which the egg has been described, do not
appear to exhibit this characteristic (Spasskii, 1951; We wish to thank Dr C. Vaucher for kindly allowing
Rego, 1961). us to examine type-specimens.
The egg of C. quadrata has a pyriform apparatus
which is attached at its narrower pole to the inner egg
membrane by a small mass of granular tissue. In whole References
eggs, it frequently appears as if the pyriform apparatus
extends at this narrower pole into elongate recurved Baer, J. G. (1927) Monographie des Cestodes de la famille des
Anoplocephalidae. Bulletin Biologique de la France et de Bel-
horns. However, if the pyriform apparatus is removed gique, Supplément, 10, 1–241.
from the egg and can be rotated under the pressure of Beveridge, I. (1978) A taxonomic revision of the genera Citto-
a cover-slip, it is apparent that the ‘horns’ are simply taenia Riehm, 1881, Ctenotaenia Railliet, 1893, Mosgovoyia
foldings of the inner membrane of the egg. It is impor- Spasskii, 1951 and Pseudocittotaenia Tenora, 1976. (Cestoda:
Anoplocephalidae). Mémoires du Muséum National d’Histoire
tant therefore that the pyriform apparatus is described Naturelle, Paris, Série A, Zoologie, 107, 1–64.
from free eggs some of which can be crushed to ex- Cabrera, A. (1961) Catálogo de los mammíferos de América del Sur.
pel internal structures, rather than from whole eggs in Revista del Museo Argentino de Ciencias Naturales ‘Bernardo
utero. Rivadavia’, 4, 1–732.
Freeman, R.S. (1949) Notes on the morphology and life cycle
The unique features of C. quadrata, with paired of the genus Monoecocestus Beddard, 1914 (Cestoda: Anoplo-
genitalia and the vagina entering the genital atrium cephalidae) from the porcupine. Journal of Parasitology, 35,
anterior to the cirrus-sac suggest that it should be ac- 605–612.
commodated in a separate genus and hence the new Joyeux, C. & Dollfus, R.-P. (1931) Sur quelques cestodes de
la collection du musée de Munich. Zoologische Jahrbücher,
genus Viscachataenia is proposed. The biogeograph- Abteilung für Systematik, Ökologie und Geographie der Tiere,
ical relationships of the new genus suggest that it is 62, 109–118.
derived from Monoecocestus, a genus parasitic pri- Led, J.E., Yannarella, F.G., Scasso, D.A. & Denegri, G.M. (1979)
Lagidiun viscaccia boxi , nuevo reservorio silvestre de Fas-
marily in South American rodents, by duplication of ciola hepatica (Linnaeus, 1758) en la República Argentina.
the genitalia. The phenomenon of duplication of the Veterinaria, 2, 31–39.
genitalia appears to be common in the Anoplocephali- Linstow, O.F.B. von (1904) Neue Helminthen. Zentralblatt für Bak-
nae (Baer, 1927; Spasskii, 1951; Beveridge, 1978). teriologie, Parasitenkunde, Infektionskrankheiten und Hygiene,
Abteilung I, 37, 678–683.
The unusual features of the eggs of Viscachataenia Mazza, S., Parodi, S. & Fiora, A. (1932) Cestode anaplocefálido
and some species of Monoecocestus further support an n. sp. de viscacha de la sierra (Lagidium tucumanus Thos.) de
association between the genera. al provincia de Jujuy. Misión de Estudios de Patalogia Regional
Argentina, 7, 1046–1054.
88

Nowak, R.M. (1999) Walker’s mammals of the world. Sixth edi- Glaucomys sabrinus (Shaw), in Oregon. Journal of Parasitology,
tion. Baltimore & London: The John Hopkins University Press, 63, 793–799.
pp. 1936. Rego, A. A. (1961) Revisão do gênero Monoecocestus Beddard,
Parra, B.E. (1953) Perutaenia thelkeldi, n.g., n. sp. (Cestoda: 1914 (Cestoda, Anoplocephalidae). Memorias do Instituto Os-
Anoplocephalidae) from Lagidium peruanum. Journal of Para- waldo Cruz, 59, 325–354.
sitology, 39, 252–255. Rowlands, I.W. (1974) Mountain viscacha. Symposium of the Zoo-
Rausch, R.L. (1976) The genera Paranoplocephala Lühe, 1910 logical Society of London, 34, 131–141.
and Anoplocephaloides Baer, 1923. Annales de Parasitologie Spasskii, A.A. (1951) Anoplocephalata. In: K.I. (ed.) [Essentials of
Humaine et Comparée, 51, 513–562. cestodology.] Vol.1. Skrjabin, Moskva: Akademiya Nauk SSSR,
Rausch, R.L. & Maser, C. (1977) Monoecocestus thomasi sp. n. 735 pp. [In Russian; English translation: Israel Program for
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